Hakan Özkan

Genetics and geography of wild cereal domestication in the near east

About 12,000 years ago, humans began the transition from hunter-gathering to a sedentary, agriculture-based society. From its origins in the Near East, farming expanded throughout Europe, Asia and Africa, together with various domesticated plants and animals. Where, how and why agriculture originated is still debated. But newer findings, on the basis of genome-wide measures of genetic similarity, have traced the origins of some domesticated cereals to wild populations of naturally occurring grasses that persist in the Near East. A better understanding of the genetic differences between wild grasses and domesticated crops adds important facets to the continuing debate on the origin of Western agriculture and the societies to which it gave rise.

Triticum dicoccoides: an important genetic resource for increasing zinc and iron concentration in modern cultivated wheat

Abstract One major strategy to increase the level of zinc (Zn) and iron (Fe) in cereal crops, is to exploit the natural genetic variation in seed concentration of these micronutrients. Genotypic variation for Zn and Fe concentration in seeds among cultivated wheat cultivars is relatively narrow and limits the options to breed wheat genotypes with high concentration and bioavailability of Zn and Fe in seed. Alternatively, wild wheat might be an important genetic resource for enhancing micronutrient concentrations in seeds of cultivated wheat. Wild wheat is widespread in diverse environments in Tarkey and other parts of the Fertile Crescent (e.g., Iran, Iraq, Lebanon, Syria, Israel, and Jordan). A large number of accessions of wild wheat and of its wild relatives were collected from the Fertile Crescent and screened for Fe and Zn concentrations as well as other mineral nutrients. Among wild wheat, the collections of wild emmer wheat, Triticum turgidum ssp. dicoccoides (825 accessions) showed impressive variation and the highest concentrations of micronutrients, significantly exceeding those of cultivated wheat. The concentrations of Zn and Fe among the dicoccoides accessions varied from 14 to 190 mg kg−1 DW for Zn and from 15 to 109 mg kg−1 DW for Fe. Also for total amount of Zn and Fe per seed, dicoccoides accessions contained very high amount of Zn (up to 7 μg per seed) and Fe (up to 3.7 μg per seed). Such high genotypic variation could not be found for phosphorus, magnesium, and sulfur. In the case of modern cultivated wheat, seed concentrations of Zn and Fe were lower and less variable when compared to wild wheat accessions. There was a highly significant positive correlation between seed concentrations of Fe and Zn. Screening different series of dicoccoides substitution lines revealed that the chromosome 6A, 611, and 5B of dicoccoides resulted in greater increase in Zn and Fe concentration when compared to their recipient parent and to other chromosome substitution lines. The results indicate that Triticum turgidum L. var. dicoccoides (wild emmer) is an important genetic resource for increasing concentration and content of Zn and Fe in modern cultivated wheat.

A reconsideration of the domestication geography of tetraploid wheats

The domestication of tetraploid wheats started from their wild progenitor Triticum dicoccoides. In this paper, the geographical distribution of this progenitor is revised to include more sampling locations. The paper is based on a collection of wild and domesticated lines (226 accessions in total) analyzed by AFLP at 169 polymorphic loci. The collection includes the 69 wild lines considered by Mori et al. (2003) in their study on chloroplast DNA haplotypes of T. dicoccoides. The goal of the experiment was to reconsider which location thought to have generated the domesticated germplasm has the highest chance of being the actual site from which wild progenitors were sampled during domestication. Phylogenetic analysis of the nuclear AFLP databases indicates that two different genetic taxa of T. dicoccoides exist, the western one, colonizing Israel, Syria, Lebanon and Jordan, and the central-eastern one, which has been frequently sampled in Turkey and rarely in Iran and Iraq. It is the central-eastern race that played the role of the progenitor of the domesticated germplasm. This is supported by the cumulative results of the AFLP data from the collections of Ozkan et al. (2002) and of Mori et al. (2003), which indicate that the Turkish Karacadag population, intermixed with some Iraq-Iran lines, has a tree topology consistent with that of the progenitor of domesticated genotypes. The Turkish Kartal population belongs genetically to the central-eastern T. dicoccoides race but at the nuclear DNA level is less related to the domesticated gene pool. A general agreement between published work on tetraploid wheat domestication emerges from these results. A disagreement is nevertheless evident at the local geographical scale; the chloroplast DNA data indicate the Kartal mountains while AFLP fingerprinting points to the Karacadag Range as the putative site of tetraploid wheat domestication.

Domestication of the Triticeae in the Fertile Crescent

About 12,000 years ago, humans began the transition from hunter-gathering to a sedentary, agriculture-based society. From its origins in the Fertile Crescent, farming expanded throughout Europe, Asia and Africa, together with various domesticated plants and animals. Where, how and why agriculture originated is still debated. Progress has been made in understanding plant domestication in the last few years. New insights were obtained mainly due to (I) the use of comprehensive germplasm collections covering the whole distribution area for each species; (II) the comparison of many wild and domesticated accessions for each species; (III) the identification of the wild progenitor in the wild gene pool and its comparison with domesticate descendants; (IV) the use of molecular fingerprinting techniques at many loci and the access to new generation high-throughput sequencing technologies; (V) the identification and cloning of genes involved in domestication; and (VI) excavation campaigns.

Evolutionary History of Wild Barley (Hordeum vulgare subsp. spontaneum) Analyzed Using Multilocus Sequence Data and Paleodistribution Modeling

Studies of Hordeum vulgare subsp. spontaneum, the wild progenitor of cultivated barley, have mostly relied on materials collected decades ago and maintained since then ex situ in germplasm repositories. We analyzed spatial genetic variation in wild barley populations collected rather recently, exploring sequence variations at seven single-copy nuclear loci, and inferred the relationships among these populations and toward the genepool of the crop. The wild barley collection covers the whole natural distribution area from the Mediterranean to Middle Asia. In contrast to earlier studies, Bayesian assignment analyses revealed three population clusters, in the Levant, Turkey, and east of Turkey, respectively. Genetic diversity was exceptionally high in the Levant, while eastern populations were depleted of private alleles. Species distribution modeling based on climate parameters and extant occurrence points of the taxon inferred suitable habitat conditions during the ice-age, particularly in the Levant and Turkey. Together with the ecologically wide range of habitats, they might contribute to structured but long-term stable populations in this region and their high genetic diversity. For recently collected individuals, Bayesian assignment to geographic clusters was generally unambiguous, but materials from genebanks often showed accessions that were not placed according to their assumed geographic origin or showed traces of introgression from cultivated barley. We assign this to gene flow among accessions during ex situ maintenance. Evolutionary studies based on such materials might therefore result in wrong conclusions regarding the history of the species or the origin and mode of domestication of the crop, depending on the accessions included.

Diversity of Macro- and Micronutrients in the Seeds of Lentil Landraces

Increasing the amount of bioavailable mineral elements in plant foods would help to improve the nutritional status of populations in developing countries. Legume seeds have the potential to provide many essential nutrients. It is important to have information on genetic variations among different lentil populations so that plant breeding programs can use new varieties in cross-breeding programs. The main objective of this study was to characterize the micro- and macronutrient concentrations of lentil landraces seeds collected from South-Eastern Turkey. We found impressive variation in the micro- and macroelement concentrations in 39 lentil landraces and 7 cultivars. We investigated the relationships of traits by correlation analysis and principal component analysis (PCA). The concentrations of several minerals, particularly Zn, were positively correlated with other minerals, suggesting that similar pathways or transporters control the uptake and transport of these minerals. Some genotypes had high mineral and protein content and potential to improve the nutritional value of cultivated lentil. Cross-breeding of numerous lentil landraces from Turkey with currently cultivated varieties could improve the levels of micro- and macronutrients of lentil and may contribute to the worldwide lentil quality breeding program.

TOLERANCE OF 65 DURUM WHEAT GENOTYPES TO ZINC DEFICIENCY IN A CALCAREOUS SOIL

Sixty-five durum wheat (Triticum turgidum ssp. durum) genotypes were grown in a zinc (Zn)-deficient calcareous soil in a greenhouse for 32 days with (+Zn=5 mg Zn kg−1 soil) and without (−Zn) Zn supply to study genotypic variation in tolerance to Zn deficiency. Genotypes were ranked for their tolerance to Zn deficiency based on the relative shoot growth (Zn efficiency ratio) defined as the ratio of shoot dry weight under Zn deficiency to the shoot dry weight under adequate Zn supply. Zinc deficiency resulted in marked decreases in shoot dry matter production of the genotypes following 32 days of growth. Shoot dry weight of most genotypes was similar under adequate Zn supply, but differed greatly under Zn deficiency. Consequently, the Zn efficiency ratio showed a large variability and ranged between 20% to 51%. Generally, the genotypes showing lower tolerance to Zn deficiency had lower absolute shoot dry weight under Zn deficiency than the genotypes having greater tolerance to Zn deficiency. Differences in tolerance to Zn deficiency were related very significantly to the total amount (content) of Zn per shoot of the genotypes, but poorly to the concentration of Zn in shoot. As the shoot Zn concentration did not vary much between the genotypes under Zn deficiency, the higher amount of Zn per shoot of the Zn-efficient genotypes was a reflection of their greater shoot dry weight. Seed Zn concentration of the genotypes was quite similar and ranged from 18 to 25 mg kg−1 DW for almost all genotypes. Therefore, the genotypic variation in tolerance to Zn deficiency was most likely inherent and not related to the seed Zn content. The results of the present study show existence of a large variation in tolerance to Zn deficiency in durum wheat, and suggest that the absolute shoot growth like the relative shoot growth (Zn efficiency ratio) can be used as a reliable plant parameter for screening the durum wheat genotypes for their tolerance to Zn deficiency, provided that genotypes have a similar growth rate under adequate Zn supply.

Diversity Assessment of Turkish Maize Landraces Based on Fluorescent Labelled SSR Markers

Landraces of maize represent a valuable genetic resource for breeding and genetic studies. Since 1970, landraces have been collected from all over Turkey, but the genetic diversity represented in this collection is still largely unknown. In this study, a sample of 98 landraces sampled from 45 provinces of Turkey was assessed genotypically at 28 simple sequence repeat (SSR) loci and phenotypically for 19 morphological traits. The landraces varied significantly for all the latter traits. A total of 172 SSR alleles were detected, giving a mean of 6.21 alleles per locus. The genetic distance between pairs of landraces ranged from 0.18 to 0.63, with a mean of 0.35. Positive and negative correlation exists among different morphological and agronomic traits. Positive association among different traits showed that improvement of one character may simultaneously improve the other desired trait. Based on UPGMA dendrogram and Neighbor-Net (NNET) analyses from both morphological traits and SSR data, respectively, it is obvious that maize landraces from the same geographical region were often placed in different clusters, indicating that grouping based on genetic parameters was not closely related to the geographic origin. The wide diversity present in Turkish maize landraces could be used as genetic resource in designing maize breeding program for developing new cultivars adapted to different geographic and climatic conditions, and may also contribute to worldwide breeding programs.

Genetic relationships among South-East Turkey wild barley populations and sampling strategies of Hordeum spontaneum

To assess the genetic diversity and the genetic structure of Turkish wild barley (Hordeum spontaneum Tell.) populations, 76 genotypes from ten ecologically and geographically different locations were analyzed by means of amplified fragment length polymorphism (AFLP) markers. Five primer combinations produced 187 scorable bands, of which 117 (62.6%) were polymorphic. Several population-specific and genotype-specific bands were identified, which differentiate populations or genotypes. Genetic distance, determined by Nei’s distance coefficient, varied from 0.07 to 0.21 with an average of 0.13. In the UPGMA dendrogram based on Nei genetic distances, the Hordeum spontaneum populations were separated into two major clusters. Genetic diversity was larger among (68%) than within (32%) populations. Eight AFLP bands were strongly correlated to the altitude of the collecting site, while no clear trend was detected between geographical origin and genetic diversity. Our results strongly suggest the need for a change in Hordeum spontaneum sampling strategy: more populations, rather then more individuals within population, should be sampled to appraise and safeguard genetic diversity in the wild barley gene pool.

Separating the wheat from the chaff – a strategy to utilize plant genetic resources from ex situ genebanks

The need for higher yielding and better-adapted crop plants for feeding the worlds rapidly growing population has raised the question of how to systematically utilize large genebank collections with their wide range of largely untouched genetic diversity. Phenotypic data that has been recorded for decades during various rounds of seed multiplication provides a rich source of information. Their usefulness has remained limited though, due to various biases induced by conservation management over time or changing environmental conditions. Here, we present a powerful procedure that permits an unbiased trait-based selection of plant samples based on such phenotypic data. Applying this technique to the wheat collection of one of the largest genebanks worldwide, we identified groups of plant samples displaying contrasting phenotypes for selected traits. As a proof of concept for our discovery pipeline, we resequenced the entire major but conserved flowering time locus Ppd-D1 in just a few such selected wheat samples – and nearly doubled the number of hitherto known alleles.